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Comparative Anatomy of the Roots in Development of Nine Epiphytes Monocots from Brazilian Atlantic Forest

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Comparative Anatomy of the Roots in Development of Nine Epiphytes Monocots from Brazilian Atlantic Forest Brazilian Journal of Development 95629 Comparative anatomy of the roots in development of nine epiphytes monocots from Brazilian Atlantic Forest Anatomia comparada das raízes em desenvolvimento de nove monocotiledôneas epífitas da Mata Atlântica Brasileira DOI:10.34117/bjdv6n12-159 Recebimento dos originais:09/11/2020 Aceitação para publicação:08/12/2020 Samara Colaoto Tessaro Doutoranda em Biologia Comparada pela Universidade Estadual de Maringá, Centro de Ciências Biológicas, Programa de Pós-Graduação em Biologia Comparada Instituição: Universidade Estadual de Maringá Endereço: Avenida Colombo, 5790, CEP: 87020-900, bloco G80, Maringá, PR, Brasil E-mail: [email protected] Rafael de Oliveira Franca Doutor em Biologia Comparada pela Universidade Estadual de Maringá, Centro de Ciências Biológicas, Programa de Pós-Graduação em Biologia Comparada Instituição: Universidade Estadual de Maringá Endereço: Avenida Colombo, 5790, CEP: 87020-900, Centro de Ciências Biológicas, Departamento de Biologia Maringá, PR, Brasil E-mail: [email protected] Káthia Socorro Mathias Mourão Doutora em Ciências Biológicas pela Universidade Estadual Paulista Júlio de Mesquita Filho Instituição: Universidade Estadual de Maringá Endereço: Avenida Colombo, 5790, CEP: 87020-900, bloco G80, Maringá, PR, Brasil E-mail: [email protected] ABSTRACT This study describes and compares root anatomical adaptations in nine monocots: Orchidaceae (Brasiliorchis chrysantha, Gomesa flexuosa, Isochilus linearis, Leptotes bicolor and Trichocentrum pumilum), Bromeliaceae (Aechmea distichantha, Billbergia nutans and Vriesea flava) and Araceae (Philodendron propinquum) epiphytes in Parana state, Brazil. Orchidaceae and Bromeliaceae have velamen radicum. Evident exodermis and endodermis in all species, cortex in the Bromeliaceae divided in three parenchyma. The immature and mature regions of the roots are different between species. Anatomical characteristics of roots related to epiphytism, such as velamen, tilosome, thick exodermis and endodermis, tracheoid idioblasts, idioblasts with raphides and aerenchyma are presents. In species that grow in the sunniest strata, it was observed that the deposition of lignin in the cell wall of the exoderm and endoderm in "O" already occurs just above the differentiated region from the primary meristems. The number of strata of the velamen was higher in these species, reaffirming that the environmental pressure throughout the evolutionary process selected genotypes that are more resistant to conditions with less water availability. Braz. J. of Develop., Curitiba, v. 6, n.12, p.9562995650 dec. 2020. ISSN 2525-8761 Brazilian Journal of Development 95630 Keywords: Araceae, Bromeliaceae, Monocots, Orchidaceae, Velamen. RESUMO Este estudo descreve e compara adaptações anatômicas radiculares em nove monocotiledôneas: Orchidaceae (Brasiliorchis chrysantha, Gomesa flexuosa, Isochilus linearis, Leptotes bicolor e Trichocentrum pumilum), Bromeliaceae (Aechmea distichantha, Billbergia nutans e Vriesea flava) e Araceae (Philodendron propinquum) epífitas do estado do Paraná, Brasil. Orchidaceae e Bromeliaceae apresentam velame. Exoderme e endoderme evidentes em todas as espécies, córtex nas Bromeliaceae divididos em três parênquimas. As regiões imaturas e maturas das raízes são diferentes entre as espécies. As características anatômicas das raízes relacionadas ao epifítismo presentes são: velame, tilossomo, exoderme e endoderme espessas, idiosblastos traqueoidais, idiosblastos com ráfides e aerênquima. Nas espécies que crescem nos estratos mais ensolarados, observou-se que a deposição de lignina na parede celular da exoderme e endoderme em "O" já ocorre pouco acima da região diferenciada a partir dos meristemas primários. O número de estratos do velame foi maior nessas espécies, reafirmando que a pressão ambiental ao longo do processo evolutivo selecionou genótipos mais resistentes a condições com menor disponibilidade hídrica. Palavras-chave: Araceae, Bromeliaceae, Monocotiledônes, Orchidaceae, Velame. 1 INTRODUCTION Epiphytes are commonly found in tropical rain forests (Kress 1986). Of the families represented by at least one habitually epiphytic species in the Neotropics, the Orchidaceae are by far the most important with ten times as many epiphytic neo-tropical species as runners-up Araceae and Bromeliaceae (Gentry & Dodson 1987). Orchidaceae comprise approximately 850 genera and 20,000 species (Dressler 1993). In Brazil, the family is represented by 250 genera and 2,760 species and is widespread throughout the country (Flora do Brasil 2020a, in construction). The species of this group have very specialized features that have allowed them to adapt to different environments; many grow in temperate regions, but they are most abundant and diverse in the tropics (Dressler 1981, 1993, Englert 2000). Bromeliaceae comprise 3,172 species and 58 genera, which are almost exclusively distributed in the Neotropics (Luther 2008). In Brazil there are 50 genera and 1,394 species (Flora do Brasil 2020b, in construction). The group is particularly suitable for epiphytic plant life, many species are endemic to certain habitats or regions, and species richness is greatest in the Atlantic Forest ecosystem (Smith & Downs 1974, Fontoura 1995, Luther 2008). Araceae comprise about 105 genera and over 3,500 species, which are distributed mainly in tropical regions and are mostly epiphytes (about 86%) (Croat 1998). In Brazil, the family is represented Braz. J. of Develop., Curitiba, v. 6, n.12, p.9562995650 dec. 2020. ISSN 2525-8761 Brazilian Journal of Development 95631 by 47 genera and 519 species distributed throughout the country (Flora do Brasil 2020c, in construction). As in other monocots, the primary root or radicle of Araceae (Coelho 2000) and Bromeliaceae (Silva & Scatena 2011) has a short life, and in the adult plant the roots are adventitious and develop from stem tissues. In orchids, undifferentiated embryos form a polarized structure called a protocorm; the posterior extremity of the protocorm forms trichomes similar to the absorbent trichomes of roots, while adventitious roots are formed endogenously in the protocorm tissue (Silva & Milaneze-Gutierre 2004b). Thus, the anatomical descriptions of the adult roots of the species in these families are based on their adventitious roots, which may have a uniseriate epidermis, with a developed absorbent hair region (Tomlinson 1969), or a pluristratified epidermis that is the velamen (Pita & Menezes 2002, Segecin & Scatena 2004, Lobo et al. 2008). The thickness of the walls of the velamen and exodermis cells varies depending on the relative air humidity (Dycus & Knudson 1957). Benzing et al. (1982) reported that the walls of most velamen cells are thin and often have perforations that allow water passage, but that the walls of cells in the inner layer of the velamen are thick. Further, over each passage cell there is the fibrous body that condenses atmospheric moisture prior to its passage to the underlying absorbing cells (Benzing et al. 1982). Pridgeon et al. (1983) called the fibrous body a tilosome. The velamen and exodermis of Orchidaceae roots have been studied extensively. Most species of Orchidaceae have velamen with or without tilosomes, an exodermis with passage cells, and endodermis cells with U-shaped wall thickenings (Pridgeon et al. 1983, Stern et al. 2004). In addition, in the majority of species the root is a polyarch and the medulla is well developed (Oliveira & Sajo 1999). According to Oliveira (2011) the cortex of aerial roots of Araceae consists of parenchyma cells and may have intercellular spaces and a sheath (sclerenchymatous ring); the endodermis, which is formed by a layer of flat or cylindrical parenchyma cells, may exhibit secondary thickening; and in the central cylinder, the xylem and phloem are alternate and surrounded by fibers or can be lobed-shaped with the xylem and phloem in an alternate and radial arrangement. The cortex of the adventitious roots of Bromeliaceae is delimited by an exodermis and endodermis, with several layers of parenchyma cells, intermediated by a sclerotic cylinder (Krauss 1949, Tomlinson 1969). Lobo et al. (2008) have described that the cortex of the adventitious root of Dyckia brevifolia and D. distachya is composed of exodermis, parenchyma, interspersed with a sclerenchymal ring, and endodermis. Braz. J. of Develop., Curitiba, v. 6, n.12, p.9562995650 dec. 2020. ISSN 2525-8761 Brazilian Journal of Development 95632 Zotz & Winkler (2013) highlighted the adaptive characters of epiphytic taxa in these families that are related to the environment in which these species grow, such as absorptive trichomes in bromeliads and the velamentous roots of aerial roots of orchids, aroids and other monocotyledons and the presence of impermeable layers that prevent loss of water after being absorbed by these structures. Given that many monocot families have epiphytic species, studies are needed to compare the anatomical adaptations of epiphytes belonging to different families native to the same location. In order to describe the anatomy of the roots of epiphytic species of the Orchidaceae, Bromeliaceae and Araceae families that occur in the same vegetation formation and whose adaptive anatomical structures have long been described in the literature, we would like to answer with this study the following questions: 1) The number of layers of the velamen and exodermis can be related to the stratum in which the species is growing most often? Species common
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